Fossil Record 9(2) (2006), 167–182 / DOI 10.1002/mmng.200600006

Baltic amber harvestman types (Arachnida: : and )

Jason A. Dunlop*

Museum fu¨ r Naturkunde der Humboldt-Universita¨t zu Berlin, Invalidenstraße 43, D-10115 Berlin, Germany

Received 15 November 2005, accepted 24 January 2006 Published online 17 July 2006

With 6 figures

Key words: Arachnida, Opiliones, Eupnoi, Dyspnoi, Baltic amber, Palaeogene, biogeography.

Abstract

Baltic amber eupnoid and dyspnoid types (Arachnida: Opiliones) in the Berendt collection are redescribed from their reposi- tory in the Museum fu¨ r Naturkunde, Berlin. Type specimens of Caddo dentipalpis (Koch & Berendt, 1854), ramiger (Koch & Berendt, 1854), Nemastoma (?) incertum Koch & Berendt, 1854, Mitostoma (?) denticulatum (Koch & Berendt, 1854) and Histricostoma (?) tuberculatum (Koch & Berendt, 1854) are all redescribed and the first photographs and camera lucida drawings of this material are presented. N. (?) incertum is removed from synonymy with M. (?) denticulatum. The status of the other Baltic amber harvestman types and their affinities are discussed. The type of bachofeni Roewer, 1939 (= S. claviger (Menge, 1854)) held in the Bavarian State collection, Munich is also redescribed here, but the repository of three other Roewer harvestman types and all of Menge’s types remains uncertain. The problematic Cheiroma- chus coriaceus Menge, 1854 is considered a nomen dubium,asisPhalangium succineum Presl, 1822, which may not even be a harvestman.

Schlu¨ sselwo¨ rter: Arachnida, Opiliones, Eupnoi, Dyspnoi, Pala¨ogen, Baltischer Bernstein, Biogeographie, Weberknechte, Spin- nentiere.

Zusammenfassung

Typenmaterial der Weberknecht-Gruppen Eupnoi und Dyspnoi (Arachnida: Opiliones) vom Baltischen Bernstein aus der Berendt-Sammlung des Museums fu¨ r Naturkunde Berlin wurde bearbeitet. Dabei wurde das Typusmaterial von Caddo denti- palpis (Koch & Berendt, 1854), Dicranopalpus ramiger (Koch & Berendt, 1854), Nemastoma (?) incertum Koch & Berendt, 1854, Mitostoma (?) denticulatum (Koch & Berendt, 1854) und Histricostoma (?) tuberculatum (Koch & Berendt, 1854) revi- diert und die ersten Fotografien und camera lucida-Zeichnungen dieses Materials hergestellt. N. (?) incertum wurde aus der Synonymie von M. (?) denticulatum herausgenommen. Der Status der anderen Weberknecht Typen aus dem Baltischen Bern- stein und ihre Stellung werden diskutiert. Sabacon bachofeni Roewer, 1939 (= S. claviger (Menge, 1854)) wird anhand des Holotypus aus der Bayerischen Staatssammlung Mu¨ nchen wiederbeschrieben. Der Aufbewahrungsort dreier weiterer Weber- knecht-Typen von Roewer und sa¨mtlicher Typen von Menge bleibt weiterhin unklar. Der problematische Cheiromachus coria- ceus Menge, 1854 wird als nomen dubium interpretiert; gleiches gilt fu¨r Phalangium succineum Presl, 1822, welcher vielleicht gar kein Weberknecht ist.

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Introduction for a review and further literature. Most of the known inclusions come from Baltic amber, which Amber is the richest source of fossil harvestmen is now usually assigned to an Eocene age (c. 44– (Arachnida: Opiliones); see Dunlop (in press) 49 Ma). Baltic harvestman inclusions were ori-

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# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 168 Dunlop, J. A., Baltic amber harvestman types ginally described by Koch & Berendt (1854), Material and methods Menge (1854) and Roewer (1939). A further, widely overlooked, name was published by Presl Koch & Berendt’s types (1822). It should also be mentioned that Berendt Berendt (1845) noted that some of the amber material he (1830) assigned some Baltic amber inclusions to was working on was in the “Mineralien-Kabinet” of Berlin. Phalangium cancroides (Linnaeus, 1758) – which In the same paper he listed species names from the forth- is in fact a Recent pseudoscorpion, currently in coming (Koch & Berendt 1854) study, rendering the 1845 names nomen nuda without diagnoses, figures or indications. the genus Chelifer Geoffroy, 1762 – and to the The “Mineralien-Kabinet” was part of the Friederich- common, Recent harvestman, Phalangium opilio Wilhelm (later the Humboldt) University in Berlin and was th Linnaeus, 1761. Although listed as an inclusion eventually combined at the end of the 19 century with the Zoologisches Museum Berlin to form the present Museum by Scudder (1891: 279), this sole example of an fu¨ r Naturkunde Berlin (MfN). Despite Petrunkevitch’s (1958, amber fossil assignable to a living harvestman p. 103) claim (at least for spiders) that the important type collections of both Koch & Berendt and Menge were lost, all species has not been substantiated in subsequent except one of Koch & Berendt’s harvestman types were work and is probably erroneous. recently confirmed as being in the MfN. Keilbach (1982) expli- Baltic amber harvestmen were reviewed by citly listed these inclusions as Berlin specimens under the unusual acronym “NPB”, specifying them as either “Original Stare˛ga (2002) based on material in Polish collec- Berendt” or, in one case, “Coll. Ku¨ hl”. The harvestmen were tions. He recognised nine valid species (Presl’s also listed under both their original and current names by name was missed), two of which he considered Spahr (1993). The Berendt fossils were formally purchased in 1873 and hard to place systematically. While Stare˛ga’s con- are associated with hand-written labels indicating the original clusions can be largely supported, he did not name and the relevant figure from the 1854 publication. make reference to the important type material of There are also repository numbers (listed below), possibly added later as they are in a different ink colour and style to Koch & Berendt (1854) held in Berlin. Examina- the rest of the label. These older numbers should not be con- tion of these types revealed that the original fused with a modern numbering scheme introduced for the authors’ illustrations are in fact reconstructions fossil collection in Berlin under the acronym MB.A. Koch & Berendt’s material is over 150 years old. The and differ, sometimes significantly, from the way amber has oxidised somewhat and is now rather dark which these name-bearing specimens appear in the ma- does not make the easy to photograph. Frequently trix. This means that subsequent authors who the amber has surface scratches or internal cracks, which occasionally obscure the inclusion. In some cases the amber have proposed transfers or synonymies (Bishop is quite brittle, fractured or even broken. Specimens were & Crosby 1924; Petrunkevitch 1955; Stare˛ga photographed using a microscope-attached Canon Power Shot G6 digital camera and processed using Adobe Photo- 1976, 2002) have done so based on somewhat shop#, and drawn with a camera lucida attachment on a Leica idealised pictures of the animals. Koch & Berendt’s MZ12 stereomicroscope. The fossils were compared to extant original material is very old and its condition is material in the Berlin collections and the literature – espe- cially Martens’ (1978) excellent summary of extant central in some cases poor (see below). Here the first European harvestmen. Familial and generic nomenclature photographs and camera lucida drawings of all follows the online catalogue of Kury (2003). the available Koch & Berendt’s harvestman type Unfortunately, the type material of one species cited to “Coll. Ku¨ hl.” – Opilio ovalis Koch & Berendt, 1854 – could species are offered as a basis for future work on not be found in a recent search of the Ku¨ hl collection in Ber- Palaeogene–Neogene amber Opiliones, along lin. This is despite the fact that Keilbach (1982) marked it with the one traceable Roewer (1939) type. with a “ þ ” in his paper to indicate that he had personally seen the specimen during a tour of museum collections in Much undescribed material exists from both Baltic 1979. The present whereabouts of the type remains unclear amber (e.g. Larsson 1978; Weitschat & Wichard and it may have been misplaced. Ku¨ hl’s amber collection was 2002, pl. 12) and the German Bitterfeld amber (see purchased by the Berlin museum in 1888 and while there are some fairly old types (e.g. mites) among it, no har- also Dunlop & Giribet 2003). vestmen were found with labels, or other indications, that The single Baltic amber laniatorid harvestman – they might be Koch & Berendt’s original material dating th a predominantly southern hemisphere clade – from the middle of the 19 century. was recently redescribed and assigned to a new (fossil) genus, Proholoscotolemon Ubick & Dun- Menge’s types lop, 2005, probably closely related to the extant Koch & Berendt’s (1854) study was posthumously published European cladonychiid Holoscotolemon Roewer, by Anton Menge from Danzig who also described additional 1915. The present paper complements this work species (Menge 1854) as footnotes to this paper. Crawford by focusing on the remaining harvestman subor- (1992, p. 15) implied that Menge’s (1854) harvestman types are also in Berlin, but they could not be found in this collec- ders recorded from Baltic amber, i.e. Eupnoi and tion. Indeed Menge’s material is cited as having been in the Dyspnoi. These include the typical ‘daddy long- Westpreußische Provinzialmuseum, Danzig (= Gdan´ sk, Po- legs’ type of harvestmen found widely across the land); see e.g. Bronn (1853–1856). After the war the Gdan´ sk material seems to have become split up. Traces of it can be Palaearctic today. found in the Museum of the Earth in Warsaw (MZW), Po-

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-fossilrecord.wiley-vch.de Fossil Record 9(2) (2006) 169 land and the Natural History Museum of Leipzig, Germany (Kosmowska-Ceranowicz 2001). These institutes might in- clude some of Menge’s arachnid types (see also below), but this has proven difficult to confirm – especially given that Menge did not provide illustrations which can be directly compared with potential type material. Note that Keilbach (1982) also included the fossil Phalangopus subtilis Menge, 1854 under Opiliones, but this taxon was listed by Petrunke- vitch (1955) as Araneae and by Scudder (1891) and Bonnet (1958) under Scytodidae (spitting spiders). It is not considered to be a harvestman here.

Roewer’s types

The types of the four species described by Roewer (1939) were part of a collection belonging to Adolf Bachofen-Echt from Mo¨ dling near Vienna, Austria. A museum repository is not given in this original paper. Keilbach (1982) cited them all as being in the Naturwissenschaftliche Sammlung des Bayerischen Staates, Munich, Germany (NSBS) – all with Bachofen-Echt collection numbers (see below). However, Keilbach but did not mark them with a “ þ ” in his list which means he did not check this repository personally. The types of Sabacon bachofeni Roewer, 1939 have been confirmed as being in the NSBS (H. Mayr, pers. comm. 2004), but unfortu- nately Roewer’s other three species could not be traced in this collection during a recent search and may not have been deposited there at all (contra Keilbach). They could not be traced in another known repository of Bachofen-Echt’s mate- rial in the Palaeontological Institute of the University of Vienna (N. Va´vra, pers. comm. 2002) either, thus the where- abouts of the three missing types remains uncertain. Stare˛ga (1976, 2002) treated all of Roewer’s species names as junior synonyms and this conclusion is supported here.

Fig. 1. A – Caddo dentipalpis (Koch & Berendt, 1854), holo- type (MfN, Berendt collection, repository nr. 7340), Baltic Systematic palaeontology amber (Palaeogene: Eocene); B – Dicranopalpus ramiger (Koch & Berendt, 1854), holotype (MfN, Berendt collection, repository nr. 7250), Baltic amber (Palaeogene: Eocene). Order Opiliones Sundevall, 1833 Suborder Hansen & Sørensen, 1904 Eupnoi 1955 Caddo dentipalpus. – Petrunkevitch: 85, fig. 53(1). Family Banks, 1893 1962 Caddo dentipalpus. – Dubinin: 481, fig. 1382. Subfamily Caddinae Banks, 1893 1975 Caddo dentipalpus. – Shear: 69. 1976 Caddo dentipalpus. – Stare˛ga: 46. R e m a r k s. The publication date of Caddidae is 1982 Platybunus dentipalpus. – Keilbach: 190. 1993 Caddo dentipalpus. – Selden: 306. sometimes given as Banks (1892), but in this ini- 1993 Platybunus dentipalpus. – Spahr: 22. tial paper only the genus Caddo Banks, 1892 was 1993 Caddo dentipalpus. – Spahr: 20. proposed. The first indication of a suprageneric 2001 Caddo dentipalpus. – Kupryjanowicz: 24, photo 3. 2002 Caddo dentipalpus. – Stare˛ga: 602, 604, fig. 2. taxon is Banks (1893: 207) who proposed a tribe H o l o t y p e. MfN, Berendt collection, repository num- Caddini. ber 7340. Ty p e l o c a l i t y a n d h o r i z o n. Baltic amber (Palaeogene, Eocene); precise locality unclear. Caddo Banks, 1892 D i a g n o s i s. Fossil caddid with three large megaspines on the ventral surface of the palpal femur and a small spinose, Caddo dentipalpus (Koch & Berendt, 1854) mesal apophysis at the distal end of the femur. Figs 1A, 2A D e s c r i p t i o n. Body oval, compact with fairly clear demarcation between carapace and dorsal 1845 Platybunus dentipalpus Berendt: 872. (nomen nudum) surface of opisthosoma. Total body length 1854 Platybunus dentipalpus Koch & Berendt: 101–102, pl. 15: fig. 125. c. 2.3 mm; maximum (dorso–ventral) thickness 1856 Platybunus dentipalpus. – Giebel: 476–477. c. 1.3 mm. Opisthosoma smooth with some hints 1885 Platybunus dentipalpus. – Scudder: 741, fig. 917. of segmental boundaries, narrowing posteriorly 1886 Platybunus dentipalpus. – Scudder: 740, fig. 934. 1891 Platybunus dentipalpus. – Scudder: 283. to form a distinct anal region. Eyes massive, 1924 Caddo dentipalpus. – Bishop & Crosby: 83–84. maximum diameter 0.58 mm, forming two lobes

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1museum-fossilrecord.wiley-vch.de 170 Dunlop, J. A., Baltic amber harvestman types which dominate the carapace region. Chelicerae to revision of Shear (1975) Caddidae is currently present, but poorly resolved. Pedipalpal femora restricted to North America, Mexico, Chile, Ja- with three prominent megaspines on ventral sur- pan, Australia, New Zealand and South Africa. face. Femur also bears mesal, spinose apophysis Like these amber inclusions, the genus Caddo is towards the distal end with short, inward-point- restricted today to the northern hemisphere. Bal- ing spines. Remaining pedipalpal articles quite tic amber shows that it used to occur more setose. Article lengths: femur 0.61 mm, patella widely, being present in north-central Europe 0.43 mm, tibia and tarsus also about the same, during the Palaeogene, but that it subsequently but boundary between them obscured. Pedipalp became extinct in this region. ends in single, gently curving claw. Legs rela- tively complete, elongate and slender, with short Family Latreille, 1802 patellae. Leg articles occasionally preserving one or two short setae. Leg 2 on left side missing. Legs often fragmentary or partly obscured mak- Dicranopalpus Doleschall, 1852 ing accurate measurements difficult, however, leg 1 c. 6 mm long in total. Dicranopalpus ramiger (Koch & Berendt, 1854) Figs 1B, 2B–C R e m a r k s. This is one of the most instantly re- cognisable amber harvestman. Bishop & Crosby 1845 Opilio ramiger Berendt: p. 872. (nomen nudum) (1924) transferred Platybunus dentipalpus Koch 1854 Opilio ramiger Koch & Berendt: 100–101, pl. 12: fig. 100. & Berendt, 1854 to Caddo Banks, 1892 based on 1854 Opilio corniger Menge: 101. the large and highly characteristic eyes. How- 1856 Opilio ramiger. – Giebel: 476. ever, it should be mentioned that juveniles of 1856 Opilio corniger. – Giebel: 476. 1891 Opilio ramiger. – Scudder: 278. other eupnoid harvestmen can have proportio- 1891 Opilio corniger. – Scudder: 277. nately large eyes and care must be taken not to 1939 Dicranopalpus palmnickensis Roewer: 1–2. confuse inclusions of young animals with caddids. 1955 Dicranopalpus ramiger. – Petrunkevitch: 86, fig. 53(2). 1962 Dicranopalpus ramiger. – Dubinin: 481, fig. 1381. The species name dentipalpus also alludes to a 1976 Dicranopalpus ramiger. – Stare˛ga: 46, fig. 79. further typical caddid character: a dentate palpal 1976 Opilio corniger. – Stare˛ga: 46. 1976 Dicranopalpus palmnickensis. – Stare˛ga: 46. femur (see Diagnosis). In not accepting this 1982 Opilio ramiger. – Keilbach: 190. transfer Keilbach (1982) appears to have been 1982 Opilio corniger. – Keilbach: 191. (as a nomen nudum) unaware of the Bishop & Crosby paper and does 1982 Dicranopalpus palmnickensis. – Keilbach: 190. 1993 Dicranopalpus ramiger. – Selden: 306. not cite it among his references. According to 1993 Dicranopalpus palmnickensis. – Selden: 306. Shear (1975) two subfamilies, Caddinae (with a 1993 Opilio ramiger. – Spahr: 22. single genus Caddo) and Acropsopilioninae 1993 Dicranopalpus ramiger. – Spahr: 21. 1993 Opilio corniger. – Spahr: 22. Roewer, 1923, can be recognised. The left palp 1993 Dicranopalpus palmnickensis. – Spahr: 21. in the holotype is overlain by an adjacent leg 2002 Dicranopalpus ramiger. – Stare˛ga: 602–604, fig. 3. which obscures the precise podomere boundary, 2002 Opilio corniger. – Stare˛ga: 603–604. 2002 Dicranopalpus palmnickensis. – Stare˛ga: 603–604. but this specimen is suggestive of an explicit H o l o t y p e. MfN, Berendt collection, repository num- character of Caddo; namely a palpal tarsus at ber 7250 (holotype of O. ramiger). Holotype of O. corniger least as long as – if not longer than – the tibia formally in Danzig (=Gdan´ sk, Poland); current whereabouts (cf. Shear 1975). Thus Bishop & Crosby’s trans- unknown. Type material of D. palmnickensis cited as NSBS (Bachofen-Echt collection, numbers 144, 149, 154); its pre- fer can be accepted. Additional material from sence could not be confirmed in a recent search. Baltic amber has recently come to light and a Ty p e l o c a l i t y a n d h o r i z o n. Baltic amber (Palaeogene, particularly fine example from the Giecewicz col- Eocene); precise locality unclear. lection in Warsaw was figured by Kupryjanowicz D i a g n o s i s. Fossil Dicranopalpus with a setose patellar (2001) and Stare˛ga (2002). There are also exam- apophsis a little over twice the length of the main body of ples of this species in the German Bitterfeld am- the patella. ber (Dunlop, unpublished observation). D e s c r i p t i o n. Body oval, compact, length Following Bishop & Crosby’s account, Shear c. 1 mm, maximum width c. 0.6 mm. Details (1975: 73) went so far as to suggest that Caddo (eyes, segmentation, ornament) of body equivo- dentipalpus is very similar to – perhaps even cal. Mouthparts equivocal. Pedipalp elongate conspecific with – the extant North American with slender articles. Pedipalpal femur length at species Caddo agilis Banks, 1892. In any case the least 0.46 mm, patellar 0.22 mm, tibia at least fossil form is of particular interest given that 0.60 mm; but distal end of tibia obscure. Patella there are no Recent records of the family in Eur- with prominent, highly setose mesal apophysis, ope, or much of Asia for that matter. According length 0.50 mm. Legs slender and elongate with

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Fig. 2. A – Caddo dentipalpis (Koch & Berendt, 1854), camera lucida drawing of the holotype; B – Dicranopalpus ramiger (Koch & Berendt, 1854), camera lucida drawing of the holotype, overview; C – detail of the D. ramiger pedipalp (fe – femur; pt – patella; ap – patellar apophysis; ti – tibia).

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1museum-fossilrecord.wiley-vch.de 172 Dunlop, J. A., Baltic amber harvestman types short patellae. Leg 2 longest, at least 13 mm long. Stare˛ga (2002), who further noted that many Leg 3 with podomere lengths: femur 1.28 mm, records are of juveniles. patella 0.36 mm, tibia 1.64 mm. Ventral surface The presence of Dicranopalpus in Baltic am- unknown. ber is notable given that this genus is predomi- R e m a r k s. This is another easily recognisable nantly found today in North Africa and southern amber harvestman species. However, the holo- Europe. As with many amber taxa, it suggest a type is actually quite a poor example which pre- previously much wider distribution. Interestingly, serves the body and legs only in outline and is one extant – albeit somewhat aberrant – species, difficult to see in the matrix. It does show the Dicranopalpus ramosus (Simon, 1909), is cur- characteristic Dicranopalpus Doleschall, 1852 rently spreading northwards through Europe; patellar apophysis (cf. Martens 1978, figs. 704–706). apparently at some speed (e.g. Hillyard 2004) However, this is essentially diagnostic for the and probably with human help. It is interesting whole (Paleogene–Recent) genus and the holo- to speculate whether these southern faunal ele- type offers little in the way of useful species ments repeatedly spread north during warmer characters. As with Caddo (see above), Keilbach time intervals – like the time of the Baltic amber (1982) either did not accept, or was unaware of, forest – only to be killed off or driven back dur- Petrunkevitch’s (1955) earlier transfer of O. ra- ing colder (e.g. glacial) periods. However most miger Koch & Berendt to Dicranopalpus. Given European Dicranopalpus species are highly spe- the unmistakable shape of the patellar apophysis cialised subalpine to alpine species with little this transfer is entirely justified. capacity for extending their range (Jochen Martens, Stare˛ga (1976, 2002) briefly synonymised both pers. comm. 2005). Opilio corniger Menge, 1854 and Dicranopalus palmnickensis Roewer, 1939 with D. ramiger.In his original description Menge (1854: 101) refer- Family ?Phalangiidae Simon, 1879 red to O. corniger as a “closely related” species – which he even admitted might simply be a Opilio ovalis Koch & Berendt, 1854 male – diagnosed by an additional short apophy- sis at the end of the pedipalpal tiba. This second 1845 Opilio ovalis Berendt: 872. (nomen nudum) 1854 Opilio ovalis Koch & Berendt: 99–100, pl. 12: fig. 99. apophysis is not shown in Koch & Berendt’s 1856 Opilio ovalis. – Giebel: 476. drawing of D. ramiger. Its absence is unsurprising 1891 Opilio ovalis. – Scudder: 277. given the poor quality of the D. ramiger holo- 1955 Opilio ovalis. – Petrunkevitch: 86. 1976 Opilio ovalis. – Stare˛ga: 46. type, which is unfortunately equivocal on this 1982 Opilio ovalis. – Keilbach: 190. character (Fig. 2C). All other figured Dicranopal- 1993 Opilio ovalis. – Selden: 306. pus material in amber (e.g. Weitschat & Wichard 1993 Opilio ovalis. – Spahr: 22. 2002, pl. 12a; Stare˛ga 2002, fig. 3) shows this sec- 2002 Opilio ovalis. – Stare˛ga: 604. ond, smaller, setose apophysis at the distal end H o l o t y p e. Cited as being in the Ku¨ hl amber collection in the MfN Berlin. Could not be traced in a recent (2005) of the tibia and given the condition of the holo- search. type its presence or absence is a poor diagnostic Ty p e l o c a l i t y a n d h o r i z o n. Baltic amber (Palaeogene, character between the first two described species. Eocene); precise locality unclear. O. corniger was listed, without explanation, by R e m a r k s. Stare˛ga (2002) regarded the systema- Keilbach (1982) as a nomen nudum, but follow- tic position of this species as uncertain. Given ing Stare˛ga it is accepted here as a junior syno- the slight unreliability of the original illustrations, nym of D. ramiger. The synonymy of D. palm- it would be unwise to comment further on the nickensis also appears justified given that affinities of this fossil until the type can be relo- Roewer (1939) described a juvenile and – as cated. with his other amber fossils – did not even men- tion the previously described species Roewer noted the characteristic (but genus-specific) apo- Family Sclerosomatidae Simon, 1879 physis on the pedipalp, yet did not figure his material and offered little in the way of a spe- Leiobunum C. L. Koch, 1839 cies-specific diagnosis. Published photographs and examination of unpublished Baltic and Bit- Leiobunum longipes Menge, 1854 terfeld material in Berlin does not suggest, thus far, the presence of more than one fossil Dicra- 1854 Leiobunum saparum Menge: 8. (? lapsus) nopalpus species in amber; see also comments in 1854 Leiobunum longipes Menge: 102.

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1856 Leiobunum longipes. – Giebel: 477. Suborder Dyspnoi Hansen & Sørensen, 1904 1891 Leiobunum longipes. – Scudder: 269. Family Dresco, 1970 1939 Liobunum inclusum Roewer: 2. 1955 Liobunum longipes. – Petrunkevitch: 86. 1982 Leiobunum longipes. – Keilbach: 190. 1982 Leiobunum inclusum. – Keilbach: 190. Sabacon Simon, 1879 1993 Liobunum saparum. – Selden: 306. 1993 Liobunum longipes. – Spahr: 20. Sabacon claviger (Menge, 1854) 1993 Liobunum inclusum. – Spahr: 20. 2002 Leiobunum longipes. – Stare˛ga: 602–604, fig. 2. Figs 3A, 4A–B 2002 Leiobunum inclusum. – Stare˛ga: 603. 1854 Nemastoma clavigerum Menge: 99. H o l o t y p e. Holotype of L. longipes probably originally in 1856 Nemastoma clavigerum. – Giebel: 475. Danzig (= Gdan´ sk, Poland), current whereabouts unknown. 1891 Nemastoma clavigerum. – Scudder: 274. Type material of L. inclusum cited as NSBS (Bachofen-Echt 1939 Sabacon bachofeni Roewer: 4–5, fig. 2. collection, numbers 51, 181); its presence could not be con- 1955 Sabacon bachofeni. – Petrunkevitch: 85. firmed in a recent search. 1976 Nemastoma clavigerum. – Stare˛ga: 46. 1976 Sabacon bachofeni. – Stare˛ga: 46. Ty p e l o c a l i t y a n d h o r i z o n. Baltic amber (Palaeogene, 1982 Nemastoma clavigerum. – Keilbach: 191. Eocene); precise locality unclear. 1982 Sabacon bachofeni. – Keilbach: 191. 1993 Nemastoma clavigerum. – Selden: 306. R e m a r k s. The original description of Leiobu- 1993 Nemastoma clavigerum. – Spahr: 21. num longipes Menge, 1854 is extremely vague 1993 Sabacon bachofeni. – Spahr: 23. and barely qualifies as an indication. Menge 2002 Sabacon claviger. – Stare˛ga: 602–603, fig. 1. 2002 Sabacon bachofeni. – Stare˛ga: 603. simply mentioned one specimen in his posses- sion under this name in which the second pair H o l o t y p e. Holotype of N. clavigerum formally in Danzig (= Gdan´ sk, Poland), current whereabouts unknown. However, of legs was (diagnostically) about twenty times Stare˛ga (2002) examined an MZW specimen (MZW 469/124) longer than the body. Furthermore, in the intro- duction to this paper he listed a different spe- cies (this time without any sort of description or indication) under the name Leiobunum sapa- rum Menge, 1854. This name was picked up by Selden (1993), but appears to be a lapsus; whereby two names (longipes and saparum) were accidentally used for a single specimen. Only L. longipes is associated with something approximating towards a description and should therefore be accepted as the valid name. It is the senior homonym of a living harvestman species; see Cokendolpher (1984) for a clarifica- tion and a discussion of Leiobunum C. L. Koch, 1839 and its unjustified amendment to Liobu- num. Stare˛ga (2002) figured only a chelicera, but commented on the fact that this species is easily recognisable and represents both the largest and most common harvestman in the Baltic amber fauna. He recognised only one Leiobunum species in Baltic amber and regarded Menge’s name as the senior synonym of Leiobunum inclusum Roewer, 1939. The type of the latter was described by Roewer as having a body length of 2.3 mm, a shiny, flat ocular tubercle lacking ornament and unmodified pedipalps with a comb-like distal claw and no apophyses on the patella and tibia. Fig. 3. A – Sabacon claviger (Koch & Berendt, 1854), holo- type of its junior synonym Sabacon bachofeni Roewer, 1939 (NSBS, Bachofen-Echt collection nr. 110), Baltic amber (Palaeogene: Eocene); B – Nemastoma (?) incertum Koch & Berendt, 1854, holotype (MfN, Berendt collection, repository nr. 7252), Baltic amber (Palaeogene: Eocene).

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Fig. 4. A – Sabacon claviger (Koch & Berendt, 1854), camera lucida drawing of the holotype of its junior synonym Sabacon bachofeni Roewer, 1939, detail; B – overview of body; C – Nemastoma (?) incertum Koch & Berendt, 1854, camera lucida drawing of the holotype.

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-fossilrecord.wiley-vch.de Fossil Record 9(2) (2006) 175 from the pre-war Gdan´ sk collection and noted that it could However, as with Dicranopalpus (see above), have been from among the material seen by Menge himself. this palpal morphology is effectively a generic Whether this material should be treated as a lecto- or neo- type requires further research into the fate of the original character. The fossil amber species can poten- Gdan´ sk collection. Type material of S. bachofeni confirmed tially be defined on its legs which, according to as being in the NSBS, Bachofen-Echt collection numbers 110 Stare˛ga (2002), are somewhat longer than in liv- (figured here) and 142. ing species (see also Figs 3A, 4A–B). Interest- Ty p e l o c a l i t y a n d h o r i z o n. Baltic amber (Palaeogene, ingly, modern Sabacon tend to occur in mountai- Eocene); precise locality unclear. nous regions today (e.g. the Alps, Pyrenees and D i a g n o s i s. Fossil Sabacon with relatively elongate legs Apennines in Europe) and it was regarded by compared to modern species, leg 2 about seven times body length. Martens (1978) as a Holarctic areal relict genus. Its presence in Baltic amber appears further D e s c r i p t i o n. Body oval, with little external north than its modern distribution. differentiation into prosoma and opisthosoma. Maximum length 2.3 mm, maximum width 1.2 mm. Carapace trapezoidal, length 0.6 mm, Family Simon, 1879 maximum width 0.9 mm. Carapace with paired, narrow lateral indents towards the anterior mar- R e m a r k s. Larsson (1978) commented on the gin and a central ocular tubercle. Opisthosoma relative frequency with which nemastomatid har- largely unsclerotised but, tergites form discrete vestmen are found in Baltic amber. Note that ovoid to quadratic sclerites which become wider the generic placements of all the fossil nemasto- and shorter posteriorly and are ornamented by matids remain equivocal based on existing type regular rows of short thorns. Chelicerae and ped- specimens. ipalps robust. Chelicerae bear numerous strong setae. Pedipalps swollen distally with characteris- tic dense setation for the genus, especially on the Nemastoma (?) C. L. Koch, 1836 tarsus. Legs elongate, fairly slender, with short setae along much of their length, femur–tibia Nemastoma (?) incertum Koch & Berendt, 1854 noticeably wider than the more distal articles. Figs 3B, 4C Leg 2 largely complete, at least 14 mm long, or c. seven times body length. Article lengths in 1845 Nemastoma incertum Berendt: 872. (nomen nudum) detail in mm. Femur 1, 1.45; femur 2, 2.45; femur 1854 Nemastoma incertum Koch & Berendt 99, pl. 17: 3, 1.36; femur 4, 2.09. Patella 1, 0.36; patel- fig. 149. 1856 Nemastoma incertum. – Giebel: 475. la 2, 0.72; patella 3, 0.63; patella 4, 0.54. Tibia 1891 Nemastoma incertum. – Scudder: 275. 1, 1.45, tibia 2, 2.63, tibia 3, 1.45; tibia 4, 2.1. Meta- 1976 Nemastoma incertum. – Stare˛ga: 46. tarsus 1, 3.63; Metatarsus 4, 3.32. Entire lengths not 1976 Mitosoma denticulatum. – Stare˛ga: 46. 1982 Nemastoma incertum. – Keilbach: 191. and/or article boundaries of more distal elements 1993 Nemastoma incertum. – Spahr: 21. equivocal. 2002 Nemastoma incertum. – Stare˛ga: 604. not 2002 Mitosoma denticulatum. – Stare˛ga: 604. R e m a r k s. The fossil described here lacks a de- H o l o t y p e. MfN, Berendt collection, repository num- velopment on the chelicerae typically seen in ber 7252. mature males of European species, although Ty p e l o c a l i t y a n d h o r i z o n. Baltic amber (Palaeogene, some East Asian males of Sabacon also lack this Eocene); precise locality unclear. protuberence (Jochen Martens, pers. comm. D i a g n o s i s. Fossil Nemastoma species lacking dorsal orna- 2005), thus it is difficult to sex this specimen un- ment of spines and tubercles. equivocally. Stare˛ga (1976) suggested that Saba- D e s c r i p t i o n. Body sub-quadrate, rounded con bachofeni Roewer, 1939 is a junior synonym anteriorly, with little external differentiation into of Nemastoma clavigerum Menge, 1854. He for- prosoma and opisthosoma. Maximum length mally synonymised them (Stare˛ga 2002) under 1.8 mm. Body widens slightly posteriorly, with a the combination Sabacon claviger; the change in distinct posterior margin. Body covered by emul- species name being associated with a change in sion, but no ornament (spines, tubercles, etc.) genus gender. Roewer’s (1939, figs 2–4) photo- protrudes through this film. Oval ocular tubercle graphs of his species are good and strongly sup- close to anterior margin of body, maximum port both this synonymy and the assignment of width 0.27 mm. Chelicerae robust, but details the species to Sabacon Simon, 1879. Both de- poor; distal articles tucked under body. Pedipalps scribed species have a highly characteristic pedi- with elongate femora, but further details poor. palp with inflated, densely setose tibiae and tarsi. Legs incomplete and partially disarticulated, but

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1museum-fossilrecord.wiley-vch.de 176 Dunlop, J. A., Baltic amber harvestman types femora of anterior legs with at least three pseu- 1993 Nemastoma denticulatum. – Selden: 306. doannulations. Disarticulated long leg article 1993 Nemastoma denticulatum. – Spahr: 21. 1993 Mitostoma denticulatum. – Spahr: 21. with distinct ornament of tiny thorns. Ventral 1993 Nemastoma succineum. – Spahr: 21. surface unknown. 2002 Mitostoma denticulatum. – Stare˛ga: 603. 2002 Nemastoma succineum. – Stare˛ga: 603. R e m a r k s. The amber piece hosting the holo- not 2002 Nemastoma incertum. – Stare˛ga: 603. type has broken. The break, unfortunately, tra- Ty p e s. MfN, Berendt collection, repository number 7250 verses the specimen through the body, but a (syntypes). Better preserved example figured here and desig- reasonable drawing of it is still possible. The nated the lectotype; other specimen (not figured) the para- lectotype. Holotype of N. succineum cited to the NSBS (Ba- holotype is also largely covered with a white chofen-Echt collection number 27), but could not be traced emulsion film which obscures some details. Star- in a recent search. e˛ga (1976) tentatively, and (2002) formally, syno- Ty p e l o c a l i t y a n d h o r i z o n. Baltic amber (Palaeogene, nymised Nemastoma incertum Koch & Berendt Eocene); precise locality unclear. with Mitosoma denticulatum (Koch & Berendt, D i a g n o s i s. Fossil Mitostoma with four rows of procurved processuli ancoriformis towards the front of the body; further 1854: see below), but did so without discussion ornament lacking. or reference to the type material. The synonymy D e s c r i p t i o n. Lectotype. Relatively complete probably harks back to Menge (1854: 99), who specimen, but missing much of the dorsal in his footnotes discussed the poor quality of opisthosoma. No external differentiation into Berendt’s fossil and found little to distinguish it prosoma and opisthosoma. Body compact, oval, from M. denticulatum. However, on close exami- total length c. 2 mm, maximum width c. 1.4 mm. nation N. incertum lacks any obvious ornament Ocular tubercle close to front end of body. Eye on the body (Figs 3B, 4C). The emulsion film is lenses small, separated by a series of small, thin enough that spines or tubercles should be visible through it. No ornament comparable to that of M. denticulatum is present and thus N. incertum can be removed from synonymy. It could either be regarded as a valid species which can potentially be recognised by its gen- eral body shape, or as a nomen dubium which is too poorly-persevered to be assigned unequi- vocally. The absence of ornament could support retaining this species in Nemastoma C. L. Koch, 1836, as in the original description, but this pre- sumably plesiomorphic feature is not explicitly diagnostic for the genus. Furthermore, the legs in the holotype appear quite long compared to typical extant Nemastoma species (Axel Scho¨n- hofer, pers. comm. 2005) – which also tend to be deep leaf-litter forms. Additional comparative material may be needed to resolve the position and status of this fossil species.

Mitostoma (?) Roewer, 1951

Mitostoma (?) denticulatum (Koch & Berendt, 1854) Figs 5A, 6A–B

1845 Nemastoma denticulatum Berendt: 872. (nomen nu- dum) 1854 Nemastoma denticulatum Koch & Berendt: 98–99, pl. 11: fig. 98. 1856 Nemastoma denticulatum. – Giebel: 475. 1891 Nemastoma denticulatum. – Scudder: 274. Fig. 5. A – Mitostoma (?) denticulatum (Koch & Berendt, 1939 Nematoma succineum Roewer: 4, fig. 1. 1854), lectotype (MfN, Berendt collection, repository 1955 Nemastoma denticulatum. – Petrunkevitch: 85. nr. 7250), Baltic amber (Palaeogene: Eocene); B – Histricos- 1976 Mitostoma denticulatum. – Stare˛ga: 45–46, fig. 77. toma tuberculatum (Koch & Berendt, 1854), holotype (MfN, 1976 Nemastoma succineum. – Stare˛ga: 46. Berendt collection, repository nr. 7248), Baltic amber (Pa- 1982 Nemastoma denticulatum. – Keilbach: 191. laeogene: Eocene).

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Fig. 6. A – Mitostoma (?) denticulatum (Koch & Berendt, 1854), camera lucida drawing of the lectotype; B – ocular region and surrounding ornament; C – Histricostoma tuberculatum (Koch & Berendt, 1854), camera lucida drawing of the holotype.

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1museum-fossilrecord.wiley-vch.de 178 Dunlop, J. A., Baltic amber harvestman types rounded tubercles. Dorsal surface largely missing to the original material. Furthermore, although from opisthosomal region, but with ornament of the dorsal surface of the opisthosoma in the lec- at least four distinctly procurved rows of proces- totype is incomplete the number of procurved suli ancoriformis, lacking any lateral connections rows of processuli is betrayed at the lateral mar- between adjacent rows. First two rows quite clo- gins and seems to have been no more than four in sely packed, c. 0.1 mm, together, third and fourth total – the ornament thus only reaching about more widely separated, c. 0.2 mm from one an- halfway down the length of the body. In described other. Chelicerae unknown. Pedipalps long and Mitostoma species there tend to be more rows slender. Distal end of femur and patella highly and this more restricted ornament in the fossil is setose with characteristic inward facing and dis- also suggestive of the probably closely-related tally globose nemastomatid setae. Legs slender, Carinostoma Kratochvı´l, 1958. Bearing in mind femora 1–3 widen distally with lengths of 0.85, the variability of structural ornament among the 1.80 and 1.11 mm respectively. Femora 1 with living Nemastomatidae, Martens (1978: 134) weak, and femur 2 with pronounced, pseudoan- preferred to separate Mitostoma and Carinostoma nulation. More distal podomeres less well pre- on genital rather than the somatic charac- served. Ventral opisthosoma segmented, but ters in Kratochvı´l’s original diagnosis. Genital details poor. characters are, thus far, not available in Baltic amber harvestmen, but Martens’ comments R e m a r k s. The original material associated should caution against raising a new (fossil) with the old label consists of two specimens, both genus based solely on different patterns of dor- in circular pieces of amber with a hole through sal ornament. the middle. The figured specimen (Figs 5A, Stare˛ga’s (1976) transfer of N. denticulatum to 6A–B) – here designated the lectotype – is quite Mitostoma, is thus tentatively accepted. This fos- well preserved, apart from the largely missing sil species is perhaps closest to Roewer’s (1951) dorsal surface of the opisthosoma. It is unclear “IV. Gruppe” whose members were defined by whether this was always the case (see below), or rows of more-or-less isolated processuli without whether the fossil has became damaged since its strong keels underlying them; a condition which original description. The other specimen – here approximates that seen in the fossil. Stare˛ga designated the paralectotype – is by comparison (1976) also referred Roewer’s Baltic amber spe- of little value, being set deep in the cracked and cies N. succineum Roewer, 1939 to M. denticula- oxidised matrix and only referable to this species tum; albeit without discussion. This Roewer type by virtue of some hints of characteristic tubercu- could not be traced (see above) and his original lation on the body. It is not figured here. backlit photograph simply shows the in Nemastoma denticulatum was transferred to outline and lacks useful details. Nevertheless its Mitostoma Roewer, 1951 by Stare˛ga (1976, assignment to Mitosoma seems justified since 2002). As in the original diagnosis of Mitostoma, even in his original description (Roewer 1939: 4) the amber fossil bears isolated rows of slightly he commented on the similarity of his amber fos- mushroom-shaped tubercles which vaguely form sil to N. chrysomelas (Hermann, 1804). This a pair of projections at their distal margins and would later become (Roewer 1951) the type spe- sometimes merge together as a continuous band: cies of his new genus Mitostoma. i.e. Roewer’s processuli ancoriformis or “Zwei- Significantly, there is no indication of spines in zackza¨hnchen” (= double-pointed teeth) or Mar- Koch & Berendt’s original description and recon- tens’ “Bru¨ ckendorn” (= bridging thorns). How- struction. They refer (p. 98) simply to “Ko¨ rnchen” ever, compared to Roewer’s (1951, figs 68–77) ornamenting the opisthosoma; albeit possibly illustrations it is worth mentioning that there are derived from this broken holotype specimen. only simple procurved rows of processuli pre- Roewer (1939) explicitly described an opistho- served behind the ocular tubercle in the fossil, soma in N. succineum without spines: “Die Tergite while many living Mitostoma species show a des Abdomens sind unbewehrt ...”. This is im- more complex morphology; specifically with a portant because there appears to be undescribed prominent row of processuli approaching, and nemastomatid material (e.g. Weitschat & traversing, the ocular tubercle. Koch & Berendt’s Wichard 2002, pl. 12f) which strongly resembles (1854, fig. 98) original drawing, reproduced by M. denticulatum in having processuli, but which Stare˛ga (1976), shows the rows being much also has a series of prominent spines in the straighter than in the actual type specimen and region of the opisthosoma tantalisingly missing in illustrates well the danger of not referring back the M. denticulatum holotype (Figs 5A, 6A–B).

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Thus there appear to be Mitostoma-like harvest- least nine in leg 4. Tarsus ends in a single curved men in Baltic amber both with and without pro- claw. minent opisthosomal spines. Provisionally, the R e m a r k s. The holotype has the typical nemas- name M. denticulatum (with N. succineum as its tomatid morphology of a body with almost com- synonym) can be used for those fossils without plete fusion of the prosoma and opisthosoma. spines towards the back end of the opisthoso- Stare˛ga’s (1976, 2002) tentative referral to the ma – even when the holotype is equivocal on extant genus Histricostoma Kratochvı´l, 1958 this character – while the undescribed spiny fos- based primarily on the paired, slender, pillar-like sils may deserve a new species name. This will spines on the opisthosoma is supported. How- have to be addressed in future papers. ever, it should be noted that slender opisthoso- mal spines can occur in other extant genera such as Mediostoma Kratochvı´l, 1958 and (the typi- Histricostoma Kratochvı´l, 1958 cally quite large) Paranemastoma Redikorzew, 1936; thus an unequivocal referral of the fossil to Histricostoma (?) tuberculatum (Koch & Berendt, Histricostoma is probably premature (Jochen 1854) Martens, pers. comm. 2005). Recent Histricostoma Figs 5B, 6C species are found in central and southern Europe, including Turkey (see also Stare˛ga 1845 Nemastoma tuberculatum Berendt: 872. (nomen nudum) 2002). One typically Alpine species can occur in 1854 Nemastoma tuberculatum Koch & Berendt: 97–98, pl. 11: fig. 97. the southern states of Germany (Bavaria, 1856 Nemastoma tuberculatum. – Giebel, 474–475. Baden-Wu¨ rttemberg) (Blick & Komposch 2004), 1891 Nemastoma tuberculatum. – Scudder: 275. but its presence here is something of an excep- 1962 Nemastoma tuberculatum. – Dubinin: 481: fig. 1379. 1976 Histricostoma (?) tuberculatum. – Stare˛ga, 45–46, tion. The fossil example from the Baltic region is fig. 78. from further north than the present day range of 1978 Nemastoma tuberculatum. – Larsson: 143, fig. 52. the genus. 1982 Nemastoma tuberculatum. – Keilbach: 191. 1993 Nemastoma tuberculatum. – Selden: 306. 1993 Nemastoma tuberculatum. – Spahr: 21. 1997 Nemastoma tuberculatum. – Krzemin´ ska & Krezemin´ - ski: 97, fig. 140. Nomina dubia 2002 Histricostoma (?) tuberculatum. – Stare˛ga: 602–603. H o l o t y p e. MfN, Berendt collection, repository num- Cheiromachus Menge, 1854 ber 7248. Ty p e l o c a l i t y a n d s t r a t u m. Baltic amber (Palaeogene, Cheiromachus coriaceus Menge, 1854 Eocene); precise locality unclear. D i a g n o s i s. Fossil Histricostoma with four pairs of small, 1854 Cheiromachus coriaceus Menge, 1854: 102. raised tubercles on the opisthosoma arranged in sub-parallel 1853–6 Cheiromachus coriaceus. – Bronn: 624. rows. 1856 Cheiromachus coriaceus. – Giebel: 477. 1891 Cheiromachus coriaceus. – Scudder: 254. D e s c r i p t i o n. A relatively complete and well- 1955 Cheiromachus coriaceus. – Petrunkevitch: 85. preserved specimen. Body more or less pear- 1976 Cheiromachus coriaceus. – Stare˛ga: 46. shaped, length 1.88 mm, maximum width 1982 Cheiromachus coriaceus. – Keilbach: 190. 1993 Cheiromachus coriaceus. – Selden: 306. 1.38 mm. No external differentiation into proso- 1993 Cheiromachus coriaceus. – Spahr: 22–23. ma and opisthosoma. Ocular tubercle present, 2002 Cheiromachus coriaceus. – Stare˛ga: 604. but poorly defined. Opisthosoma bears four pairs H o l o t y p e. Formally in Danzig (= Gdan´ sk, Poland), current of short, blunt dorsal spines, forming two rows whereabouts unknown. c. 0.2 mm apart. Chelicerae present, but largely Ty p e l o c a l i t y a n d h o r i z o n. Baltic amber (Palaeogene, indistinct. Pedipalps with elongate femora bear- Eocene); precise locality unclear. ing numerous, mostly inward-facing, setae. More R e m a r k s. This species is significant as the only distal articles hidden within the matrix. Legs Baltic amber harvestman originally assigned to more or less complete, slender and still articu- an extinct genus. While the Baltic amber lania- lated to the body. Legs 1–4 with approximate torid, P. nemastomoides, differs sufficiently from lengths of 4.6, 9.6, 4.9 and 7.5 mm respectively, living taxa to warrant a new, fossil genus (Ubick i.e. a leg formula of (from longest to shortest): & Dunlop 2005), all other Baltic harvestmen 2, 4, 3, 1. Leg articles with occasional short have thus far been assigned to extant genera. In setae. Leg 1 femora with possible hints of pseudo- the footnotes to the original description of articulation. Tarsi divided distally into multiple P. nemastomoides, Menge (1854) suggested that tarsomeres (not resolvable in all legs) with at this laniatorid should effectively be placed in a

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1museum-fossilrecord.wiley-vch.de 180 Dunlop, J. A., Baltic amber harvestman types different suborder (Eupnoi) via the genus rial thus remains unclear and is probably lost (Wolfgang Acantholophus C. L. Koch, 1839 (preoccupied, Weitschat, pers. comm. 2005). Presl’s work incidentally is the oldest historical record I am aware of offering formal de- now Lacinius Thorell, 1876 – see discussion in scriptions and names of fossil . It is thirteen years Crawford 1992). Later (Menge 1856: 11), he older than the Coal Measures scorpion Cyclopthalmus senior changed his mind and suggested that P. nemasto- Corda, 1835 which Petrunkevitch (1953: 24), for example, cited as the (historically) oldest fossil arachnid to have been moides was “probably identical” to Cheiroma- described. chus coriaceus Menge, 1854, although without an Ty p e l o c a l i t y a n d h o r i z o n. Baltic amber (Palaeogene, illustration of the latter species this assignment is Eocene); precise locality unclear. difficult to test. Stare˛ga (1976, 2002) regarded R e m a r k s. Apart from being listed in Scudder the systematic position of Cheiromachus as un- (1891) the name has been overlooked by all sub- certain within the Phalangiidae. In an attempt to sequent authors. The original Latin description resolve this situation, Menge’s (1854: 102) origi- of a c. 4 mm long animal is very general and nal diagnosis – reproduced by Bronn (1853–6) could apply to any number of taxa. No figure and as a shorter English translation by Petrunke- was included. The fact that Presl described long, vitch (1955) – is retranslated here as follows. dark legs, “Pedis fusci, longi ...”, a quadratic “A new genus Cheiromachus distinguished by prosoma, “Thorax quadrangulus ...”, and – sig- peculiar pedipalps: the first article short and in- nificantly as a separate body region – an elon- versely conical. The second article very thickly gate, oval opisthosoma, “Abdomen oblongum, curved backwards, laterally somewhat pressed ovoideum...”, raises the possibility that Presl together, at the lower surface pustulate, rather mistook a spider for a harvestman. Opiliones long. The third article inversely conical, much generally do not have an elongate opisthosoma thinner and half as long. The fourth is short and and the basic body shape he described would be spindle-like. The final article thin and wiry. The consistent with certain spiders like, for instance, end, however, is missing. Likewise all limbs up the long-legged Pholcus which is sometimes con- to some of the thighs missing. Body complete. fused with harvestmen by the general public. Clearly visible is the pustulate dorsal surface of Without a specimen or illustration with which to the body. The pustules on the raised, almost confirm the identity and affinities of this taxon, wart-like ocular tubercle stronger, the ventral Phalangium succineum Presl, 1822 should prob- 000 surface smooth. Length over 2 . Abdomen ably be regarded as a nomen dubium. width 1000. Ch. coriaceus.M.” A number of extant genera exhibit a compar- ably pustulate body and/or ocular tubercle, but these ‘peculiar’ thickenings on the pedipalps de- Acknowledgements scribed by Menge cannot be readily matched to any known (living) genus (Jochen Martens, pers. I thank Christian Neumann (Berlin) and Helmut Mayr (Mu- nich) for access to material in their care and Axel Scho¨ nho- comm. 2005). Conceivably there is an extinct am- fer (Mainz) and Plamen Mitov (Sofia) for valuable comments ber harvestman genus with diagnostic pedipalps on harvestman . Barbara Kosmowska-Ceranowicz and future work may recover specimens compar- (Warsaw), Norbert Va´vra (Vienna), Vojteˇch Turek (Prague) and Wolfgang Weitschat (Hamburg) kindly offered informa- able to Menge’s description. However, in the tion about known or likely type repositories. Jochen Martens absence of a type specimen or illustration it is (Mainz) and Christian Komposch (Graz) made numerous probably best to place Cheiromachus coriaceus helpful observations, both through correspondence and their reviews of the manuscript. for the time being as a nomen dubium of uncer- tain familial affinity.

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Bishop, S. C. & Crosby, C. R. 1924. A fossil species of Caddo Kratochvı´l, J. 1958. Ho¨ hlenweberknechte Bulgariens (Palpa- (Opiliones) from the Baltic amber, and its living rela- tores – Nemastomatidae). – Acta Academiae Scientiarum tives. – Bulletin of the New York State Museum 253: Cˇ echoslovenicae Basis Brunensis 30: 523–576. 83–84. Krzemin´ ska, E. & Krezemin´ ski, W. 1997. Les Fantomes de Blick, T. & Komposch, C. 2004. Checkliste der Weberknechte l’Ambre. Insectes fossiles dans l’ambre de la Baltique. Mittel- und Westeuropas./Checklist of the harvestmen of 142 pp., Muse´e d’Histoire naturelle de Neuchaˆtel, Suisse, Central and Western Europe (Arachnida: Opiliones). – Neuchaˆtel. Internet: http://www.arages.de/files/checklist2002opilio- Kury, A. 2003. Checklist of the valid opilionid genera of the nes.pdf. 6 pp. world. Bonnet, P. 1958. Bibliographia Araneorum, Tome II (4me http://acd.ufrj.br/mndi/Aracnologia/checklaniator.htm partie N-S). pp. 3027–4230. Douladoure, Toulouse. 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